A STUDY OF As?lenium platyneuron (L.) Oakes
AND
Camptosorus rhizophyllus (L.) Link
\-l1TH AN .SllPH..t::..2IS
em
SPOiC:':
10Rl.~HOLOGY
A Senior Paper
Submitted to
Or. J. C. r.falayer
of
Ball State University
by
Lois A. I(inder
In Partial Fulfillment of the Requiren.ents
for graduation
on
The Honors Program
l'.J:ay I, 1966
:;;rCo~1
7he:!
ii
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TABLE OF CONTENTS
Page
LIST OF TABLES ••••••••••.••••.••••••••••••••.••..•
......... .. . .. ..............
INTRODUCTION •••••••••••••••• ............... . . . ....
LIST OF ILLUSTRATIONS.
iii
iv
1
LI Ir .2.PJ.l.TUl<'E ••••••••••••••••••••••••••
2
Morphology ••••.•••••..••••••••••••••••••••••••••
2
Taxonomic Realtionships •• •••••••••••••••••••••••
4
REVIE\~
OF
irH~~
NETrIODS
MATERlii.LS ••
............... .... .. . . ....
6
General 1>1orphology ••••••••••••••••••••••••••••••
6
:-")pore }forphology ••••••••••••••••••••••••••••••••
7
. . . ... . . . . . ... . . . . .. . . . . . . . ..
General Horphology..............................
Spore Morphology................................
'[lATA •••••••••••••••••
DISCUSSION.
... . ... ................................
15
15
30
39
sm~J~y...........................................
46
BIBLIOGRAPHY......................................
47
iii
LIST OF TABLES
Page
Table
I.
II.
Gross morphological calculations
of Gamptosorus rhizophyllus (L.) Link
I. A Specimen measurements •••••••••••••••••
8
I. B Leaf height analysis ••••••••••••••••••
19
Gross morphological calculations
of iisplenium platyneuron (L.) C.akes
II.
III.
Specimen measurements.................
9
II. B Leaf height analysis •••••••••••••••••
24
A
:}ross morphological calculations
of Asnlenium
ebenoides (Scott) Wherry
+
III. A
measllrenents................
10
III. B Leaf height analysis.................
28
III. C
IV.
S~ecimen
i~ea
variation measurements..........
29
Spore size analysis of
Camptosorus rhizophyllus •••••••••••••••••••
12
IV. A Spore length
32
IV. B Spore "7idth
v.
VI.
•••••••••••••••••••••••••
.........................
33
Snore size analvsis of
Asplenium platyneuron ••••••••••••••••••••••
13
V. A Spore length ••••••••••••••••••••••••••
34
V. B Spore width •••••••••••••••••••••••••••
35
Spore size analysis of
..:\splenosorus ebenoide.s (Scott) vJherry ••••••
14
VI. A Spore
•••••••••••••••••••••••••
36
VI. B Spore width ••••••••••••••••••••••••••
37
len~th
iv
LIST OF ILLUSTRATIONS
Figure
Page
Mature sporophyte of Camptosorus
rhizophyllus (L.) Link ••••••••••••••••••••
17
llature sporophyte of Asplenium
platyneuron (L.) C'al:es ••••••••••••••••••••
21
Nature sporophyte of .I:\splenosorus
ebenoides (Scott) r,jherry ••••••••••••••••••
26
4.
Photograph of C. rhizophyllp.s •••••••••••••
18
5.
Photograph of A. platlIleuron ••••••••••••••
22
6.
Photograph of A. ebenoidew ••••••••••••••••
27
7.
Spore material of
rhizophyllus •••••••••
42
8.
Spore naterial of
.(i.
platyneuron ••••••••••
43
9.
Spore material of
~.
ebenoides ••••••••••••
44
1.
2.
3.
INTRCDUCTIGH
'.Chis study 17as made in order to learn about substantiating a possible taxonot:lic reJa tionshiT> b'~b"ee.n
,'~splenium
platyneuron (L.)
rhizophyllns (L.) Linl:.
O[ll~es
'l'hrou\:,;h
and Camptosorus.
acquaintance \·yi th
all.
methods 0:'::: spore analysis em,; simple slide prep;lration,
it nas ~)ossible to [,lake observations
0;,'
the Gross
rn.orpholo?,;Y 8n<1 the spore 'lorpholo::::s \·"hich these
:Cern species possess.
hlO
As a result of the kind coop-
eration of the Indiana University T-IerbarillD1 an.d the
Chica30 Natural History Euseum it was )ossihle to examine borrm,;red
.
sDecirl~ns
.
of Asnlrmium ebenoide.s (Scott)
T::herry, a reported hybrid o£
rhizophyllus.
--~.-------.!l..
----------
rJlntyneuron and ._.
In addi ti.,n to indirect studies, :::m
atte;'1pt '\Jas Tl:;de to
~
~:~roH S')ecirl~ns
_ _ . . L .
1)£ both,\.
,)lEl't:'vru~uron
_ _
_ _
y~~
_ __
"mel _G. r~d.zoi)h·.J.lllus.
e _
.. _
By applyin;;:
"-'" t:re(~ cliffe ·ce.nt: ,'le~chods
of cultnrin,: the fern
S~)ore.s
, it va:;; hoped tlLa:i: suf:i:icie.nt
r:n terial cDuld be obtained f'')r c08parative
c:mnts
0:(
the. ferns.
chrOi~tOSOile
2
.i:U!:VIE\·J Gl" '1'1-1£ LI'i'EJJl.'I'URE
In the study of -Cern taxOnCY1Y, spore exar,l:lnation
haf3 an important function,
'}'11e ch.aracteristics of color,
size, shape and structural f'13rldnt;s are used :':or id'~nt-
-.c-
4-
~£~caL.~on.
•
Spore m'::lr:1holor;Y lS stron:;;ly supported os a
means of: identification be.cau.se it can help to c~istin;uish ·t::he species of one c;enErll; differen:i:iate genera;
and characterize :Cflnilies.(3)
that anatomical
8J~
fern are valuable.
Some Horkers also feel
cytological studies of
~he
entire
In .9.ddition to indirect studies, airect
tests ;':'l1d experinents are hoped to reveal i:urther
ledge concerning fern taxonomy.(16)
~moH-
In future studies it
ruay be possible to better identify 901yploid ntrains of
kno1;VIl species as ,(·:cll as to indicL:te hybridity.
Nost of the p:2St studies
:~roupings
involvin:~
the
~::,~~p1(~nium
have involved norphology, anatomy, and
cytolo~y.
Studies conducted by 8. '1'. "'herry during 1925 and 19.36
di vided the Hhole Asplenium ;>;roup into three original
species of extreme variation.
i.
':i..
nont8num.., A. pl.atyneuron,
rhizophyllum (Q. ;rhizophyllus, Linl<) ·'?!re the
l~;ostulated
ancestral types vlhich joined by A. pinnatifidu!'.1 and
~.
bradleyi "vould compose five basic species of the srouping.
Dr. tJherry t s direct evidence cf ·this
h~7:)othesis
came from
3
the production of
the postulated forms by hybridi~ation
experiments Hhereb~T eleven describuI entities ~'lere proIn 1953 D. H. Britton published Em article
duce.d.(16)
of his 1'!ork on chrol'losome. studies of ferns.
t;u:,~~~ested
It\·JaG there
th.E:t Phyllitis scolo)?endrium var. j:'meric[:lna
Fern, collected in Durham, Ontario, could be considered
closely related to Camptosorus rhizophyllus (L.) Link
according to the 93metic chro11Osome tllUuber. (2)
of U.
t-~.
1:'.;,
study
Uagner conducted at approximately the SHrle time
concern~d
1:'{,e species and hybrids centering about
montanum, J\. qradleyi, ;".
pinn~~fidum.
(15)
H.
Int:E~rmed-
iate series of these thre.e species ,'Tere. illustrated by the
1;vork of H. D. Gray Clnd Tv. T. Vherry.
neuron and
"l
t.-, •
Including
ll..
platy:-
rhizophyllus fuuong the h.ppalacLi.ian As;)len-
iums, the follo\r.i.ng hybrids Here suggested:
A. mont anum x
c.
rhizophyllus - A. pinnatifidum
A. montanum x A. platyneuron -
~.
bradleyi
A. ;>latyneuron x G. rhizophyllus - X Asnlenosorus
~benoides(15)
Further studies of Va::sner \!ith Kathryn Boydston in 1958
indicated three basic species of the Appalachian Aspleniums as A. T."10ntanum, A. platyneuron, and A. rhizophyllus.
These in turn 'ive.re. thought to generate nine taxa one of
'Iahich has t'\-70 cytological forms.(17)
.ii
chromatographic
4
study of the "'>.ppalachian Asple.nium cor:ple~: H~IS published
in 1963 by Dale L. Smith and Donald A. Levin.
All results
of their study re.ve.'~led a confirmation of \'Ya:;:::ner l s ,,!ork
(1954) accordin:::: to th:::: conparative norpholo:S'y, hybridization, and '::aryology.(10)
A :positive hypothe.sis of
direct evidence nmv supported ;)ostulc1tef> m::,de. about the
interrelationships in the )I.splenium Group.
Several Horkers in fern taxonomy, among them Hagner
Ivwing published several papers, believe As;)lenium ebenoides
to be the
v[~lid
hybrid")f AS2lenium platynelU'on and IJamp-
tosorus rhizophyllus.
speClE~S
in 1865.
;{. R. Scott decL.,.red i.t a ne'V7 fern
N. J. BerIdey in 1366 declared A. ebenoides
to be the natural hybrid of A.
~)latyneuron
x
c.
rhizophyllus.
Some persons violently objected to this cOi1clusion; rmd
the question
:)f
h~'bridity ,'las
finally resolved by lIargaret
Slosson in 1902 durinc; the course of her experiments of
artificial spore cultures.(13,12)
Early specimens of the
hybrid Here single, sterile plants, but a
fertilE~
popula-
tion was discove.red in Havana Glen, Alabama prior to 11iss
Slosson's \York.
In 1910 H. D. Hoyt sought to disprove the
existence of "1:1.1e hybrid for the evidence seemed insufficient
to him.(l8)
A sterile diploid popUlation exhibiting vDried
growth stages 'i;·!as rliscovere.d by ivagner in 1946 in
Nontgomery Co., lv!.qryland. (11)
5
Further studies conducted in 1953
~7ere
the first of a.
cytological nature ,:)n the Alabama ;Jopulation.
~'7ere f·~und
sterile ·qnd ::ertile hybrids
Hork concerning the reL:'.ti')l1ship of the
lns
prc>sress:~d
Both
to occur. (18)
;:~s0l(~nil1.m
2:roup
from an indirect procedure involving
color, size, and structural markings to a more 6.irect
procedure involving artificial cultures, chromosome
counts, ;mel biocll .emic-31 consti tue.nts.
i\sDL~.niupl
~
discover
Knowle.dge. of the
habitat hag also ')rn:1ressed
as v."rious \vorkers
.......
ne~"
-
populBtions of the basic species and their
reported hybrids.
6
HETHODS
L.im
i\A 'fERI.AlB
General Norphology
During the SUrUl::ler of 1965 specim2.ns ef ,:lsp}.enium
(~amptosorus
platyneuron (L.) Oakes nn":
rhizophyllus
(L.) Link ,lere collected in Gre::ne, Eonroe, and Parke
counties of IndL.ma.
The dried spc.cir,1ens we.re. Lien cxam-
during SeJ)tember thro1J:'::h February Df the follO\"i.ni:~ months.
in addition to these recent specin::!ns of A. pL::tyneuron
and
c.
rhizophyllus used for study, olde.r
s'~)ecimens
of
A. ebenoides 'Nere borroHed fre:D.1 the Indiana Univen.lity
Herbarium gnd the
ehica~;o
Natural i'istory l':useum.
Struc-
tural m2.asurements ,·Jere taken of several rlants and size
variatic:n s ,,,ere computed.
Snecimens
uere coll:-;cted
for the Flost ;;:lrt in
"-
~
moistened plastic bags and then transferred to a plant
press.
lTnole plants including the rootstocks Here care.-
fully dug
fr~)m
their sites.
be washed or shake.n m..,ray.
mntent of tl'.e ferns,
Before dry ins ,the dirt could
De;::>endinr; upon the 80isture
the dryin2: process could
pleted Hi thin one to t"l'70 ,';reeks.
b0..tl,'leen sheets
O~~
ne'ivS 1)8pe.r
blotter "9 arti tinns.
site and date at
t~ie
bE~
com-
The Dlants were dried
I,!i thin
a vJOoden pre.ss of
SpeciL1ens were labeled as to the
tL'le of collection.
PlAnts 'ivhich
7
previously had be;~n dried Her:: stored inside a box l)et~veen
lflbeled sheets
0
f newsJ:"l8per.
Later
-;:1'0
se specin~ns desired
for flJ_ture reference and spore s]mples
~,rere
stron~~
Struetur<ll T'Jcasure-
constrllcti,m paper ;:md st,)red.
ments :)f A. platyneuron,
T!7ere made
ryf
,j.
rhizophyllus,
<l.
fllounted on
ebenoide~
fromd len-;th, stem length, and Greatest frond
~'7idth usin~ D 15 em ruler. (See Fig. I-VI; Tables 1 A-3 A)
Spore Horphology
L4 rorn -the dried material of C. rhizophyllus and
l'i.
platyneuron, spores 1;12re obtained for simple clearing
and staining techniques.
r:1ountin~~
'1'he first slides l'lade :;_nvol ved
a :C"eiv dried s:)ores nn clean slides
Ined:i_um; rmd Glycerine jelly
IV~AtedJver
To ei the-c f,10untin[': solution
-:1
Clear nail
e,:,ntrast stc-:in of
factory sealer;
-~)olish
I-Ioyer f s
an alcohol lamp.
Vethylene Blue::r ?ast Bre2n was then added
slip applied.
usin~~
3'(1(1
t~ither
Cl
cover
was f{)lJ,nd to be a aatis-
:nd slid2S Here stored flat for :final
.lrying and st'Jrag:;.
In ele,"lrin'c; th(O'. fern
spol~es,
hydrochloric , c::c2tic,
and suli)lmrin acids Here used as ;:.:zents.
proved [lost satisfaetury
bOt~1
fron tile standpoint: ::Jf
tile also11ol flame.
:,]aution T·ms nec0.sSf1!ry to avoid
scorchinr~
Fl'-lterial.
(J
the
exceed CO C.
s~')or~
Tenper;-lture [;hould not
8
Table I
.f!.
Specimen n -:,asure':ne.nts of
Camptosorus rhizophyllus (L.) Link
Collection
Site
?nrke Co. ,
Indi2na
"
II
Leaf Height
8.0
19.5
28.5
27.2
9.7
14.0
26.0
18.5
17.1
11.0
17.1
9.5
23.0
14.7
11.3
9.5
8.8
6.2
17.7
29.5
21.4
4.9
14.5
6.0
CIll
cm
cm
Frond
Stem
6.7
18.0
24.5
23.0
5.2
10.3
23.0
15.8
13.9
5.5
14.1
8.0
18.0
11.7
8.7
6.0
7.0
3.3
15.5
1.3
1.5
4.0
2~.5
7.0
6.0
1.0
1.2
1.5
\.Jidth
1.0
1.2
1.6
1.7
1.6
0.7
1.6
1.8
1.7
1.4
1.3
0.9
1.5
0.9
0.9
1.5
0.7
1.2
0.7
2.0
2.0
1.0
1.6
1.1
Total:
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
391.1 em
306.1
72.0
32.2
Hean:
15.6 em
12.2
2.9
1.3
II
"
"
fI
II
11
H
"
II
II
"
II
"
It
If
It
"tt
"
tt
If
15.4
3.9
13.3
4.5
l}.2
L;-.5
1.0
3.0
2.7
3.2
5.5
3.0
1.5
5.0
2.0
2.0
3.5
1.8
2 a
.~
2~2
9
Table II A
Specimen measurements of Asplenium
Collection
Site
Leaf Height
Greene Co. ,
Indiana
16.5
28.8
31.7
32.0
19.5
12.0
15.0
29.0
25.7
17.0
17.5
lS.O
16.0
19.5
12.5
25.2
20.8
22.8
13.7
15.6
"
"
"
ronroe Co.
Greene Co.
"
Co.
Co.
Co.
Co.
Co.
Tf
l~ean:
em
Hidth
2.0
3.9
3.7
3.0
5.0
2.5
3.5
5.5
3.3
2.0
3.0
2.5
3.0
4.0
1.5
2.8
2.5
2.4
342.2
63.4
50.9
17.1
3.2
2.6
20.3 em
fJ
Stem
(L.) Oakes
2.5
3.2
3.5
2.3
2.5
1.7
2.7
2.8
2.0
2.2
2.5
3.0
2.3
3.3
2.4
3.1
2.0
2.4
2.1
2.4
2~.9
Total: 405.8 em
Co.
"
"
Nonroe
Greene
Honroe
Greene
}fonroe
14.5
em
em
em
em
em
em
em
em
00.
If
Green~
Frond
28.0
29.;0
14.5
0.5
ll.5
23.5
22.4
15.0
14.5
12.5
13.0
15.5
11.0
21.2
17.5
2Q.0
11.2
13.2
II
Nonro~
em
em
em
em
em
em
em
em
em
em
em
p18t~euron
1+.0
"
..
_J. J
10
Table III A
Specimen measurements of Asplenium ebenoides (Scott) ;,inerry
Collection
Site
Co. ,
Indiana
Uilmington,
Delat,rare
~'1onroe
II
II
College Hill
Easton, Pa.
"
It
Havana Glen,
Hale Co. , lila.
II
II
If
"
If
II
rl
II
If
Slos.on
culture
"
If
II
II
If
If
II
II
ff
Leaf Height
15.5
18.0
3.4
7.3
3.6
6.6
5.0
4.3
10.7
14.8
15.0
14.0
7.0
12.5
5.5
6.4
14.7
11.8
11.6
13.5
14.4
10.3
9.6
11.7
11.8
11.9
15.5
12.3
13.3
11.0
G.O
9.4
em
em
em
em
em
em
em
eP.l
ern
ern
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
Frond
11.0
12.5
1.3
3.1
2.3
4.6
l~.O
3.4
8.7
10.8
6.0
5.5
3.8
9.5
3.2
5.0
8.7
6.8
6.8
7 .L~
11.4
9.5
3.7
9.7
11.0
9.5
13.5
9.3
11.2
10.3
6.3
8.8
Hidth
Stem
4.5
5.5
2.1
4.2
1.3
2.0
1.0
0.8
2.0
4.0
9.0
8.5
3.2
3.0
2.3
1.4
6.0
5.0
4.8
6.3
3.0
0.8
.
o
<..)
."
2.0
0.8
2.4
2.0
3.0
2.1
0.7
1.7
0.6
2.7
2.3
1.0
1.8
1.1
1.8
1.0
0.8
4.0
t::
...
-,.:J
2.0
2.0
1.2
1.3
0.4
1.0
2.5
3.0
1.9
2.0
2.3
2.0
2.1
1.9
1.8
1.9
3.6
3.2
2.8
1.7
2.0
1.1
11
The acid Has decanted ;-nd the
s~)ore
material then rinsed
t'Y70 or three time.s ,'lith distilled i'later.
decanted and s small sample of the
a clean slide.
The \Tater
s~)ores
\'laS
the.n placed on
Lost of the I,later ';1as allO'iled to eVapDrate.
Both Hoyer's and Glyc,'>.rine jelly Here selected as muunting
medias; 2nd the ss.me. C"_nlTasting stains of IJethylene Blue
and Fast Green used.
st~lred
flat :enr (1.rying.
ed. by Zrdtoan
1:!8S
:,c:,n acet:'lysis ;'Jroceclure
2.S
explain-
c('!nsidered (5); but yr:Ned unnecessary
to obtain clearing results desireable for this observ2.tion.
The t'OvO stains used for slight color contrast 'Here not
thought to be essential for an accurate exarlination of the
spores
'>Jhe:i..~e
the clearin::; tec0niqucs I'7ere succe.ssful.
The follm'7ing Figu?:es'7,8,9 illustrate the results cf spore
observations concerning
-
and A. ebenoides.
c.
rhizophyllus,
!!.
platyneuron,
Haterial from the latter sDecimen \las
-
0bserved in a simple slide prenaration nsing Hoyer's medium.
The othe.r slides i,!ere mnde of beth Hoyer's and the Glyce.rine
jelly.
lens.
All spore draHings 'hrere made lmder an oil iDmersion
Neasureme.nts I-Jere made \'li th an ocular nicrome.ter
wherein ei:?;ht snaces of the eyepiece equaled O.Olmm at 95x.
(See Tables 4 thr:)ugh 6)
12
Table IV
Spore Size Analysis of Camptosorus rhizophyllus (L.) Link
NeasureI'1ents
Collectioh
Site
Parke Co.
Indiana
If
If
If
tI
fT
fl
ff
If
II
tI
II
II
II
II
II
If
fI
If
II
If
IT
tt
tI
tI
tf
tI
Length
34
20
26
20
37
34
40
26
31
'23
>7
40
34
31
43
40
31
34
43
31
28
30
26
37
26
34
Hidth
fiU
fiU
mu
rou
rou
fiU
rou
mu
mu
mu
mu
mu
rou
mu
mu
mu
mu
mu
fiU
rou
mu
fiU
fiU
mu
mu
fiU
28 mu
28 mu
ll~
')
,;:
...
-,..
l--:lU~
fin
lL:~
r:.m.
23
r:1U
28
20
20
14
20
28
23
26
31
28
20
31
28
26
31
31
28
31
17
31
mu
mu
mu
mu
mu
mu
mu
mu
mu
mu
mu
mu
mu
r,
2 (,
nu
fiU
rou
fiU
nu
rou
rou
rou
23 rou
13
Table V
Spore lvIeasurements of Asplenium p1atyneuron (L.) Cakes
Collection
Site
Greene Co.
Indiant
II
r:'idth
Length
:54 mu
31 rou
37 rou
mu
II
::)4
If
.20 QU
II
30 mu
37 mu
54 mu
31 fiU
}\,-!nroe Co.
Inditlna
If
ff
L~O
ff
28
26
28
37
40
34
31
II
II
t!
"
II
"ff
n
It
ff
II
"
"
TI
28
37
40
28
34
43
28
:54
mu
mu
mu
rou
rou
mu
mu
mu
mu
mu
mu
mu
mu
mu
mu
mu
31
22
27
21
10
20
28
mu.
mu.
mu.
E1U
mu
rou
mu
"_u mu
281 mu
28 mu
')0
')oop
.c..J fiU
20
23
:A
26
26
20
23
:6
)6
17
mu
mu
mu
mu
mu
mu
nu
fiU
mu
mu
28 mu
31 mu
"'_J nu
26 mu
')~
L4
'l'able VI
3pore. lIeaS1.lreme.nts of
J:~~sElenosorus
Col1eEtion
Site
Length
1'onrne Co.
Indiana
II
fI
It
"
It
It
If
It
It
"
II
II
Hidth
20 mu
10 r.Ill
15 mu
16 mu
[3 mu
14 mu
10 mu
7 mu
11
12 r.,lU
7 mu
'~'lU
5 1.:lU
6 mu
"
14 mu
II
8
;'lU
II
11
17
11
14
20
mu
mu
mu
mu
mu
3
P1U
I'
"
"
"
"
II
II
I
r
II
ebenoides (Soott)
14 mu
20 faU
14 ruu
10 mu
17
7
13
13
7
12
mu
mu
r,l.U
I~lU
m.u
Ei.u
8 nu
6 nu
10 nu
1.1 mu
6 r.lU
3 mu
6 nu
n
mu
0
6 mu
CJ
" mu
14 nu
6 mu
11 mu
14 mu
6 mu
11 ElU
11 mu
12 mu
6 mu
~·Jh.erry
15
DATA
gene.ral l i orpholor;;y
'fhe majority uf fern specimens observed for this
study ':'lere ccllected in the Indiana ccmnties of Greene,
Honroe, and
P.~rke
during the summer of 1965.
Specimens
uere also borr'.:.med from the Indiana Dnivel'sity llerbarium
and the ChiCago Natural History Nuseum.
F()r these spec-
imens observed gross structural measurements and spore
measurements 1;Iere taken: and size variations Here computed in order to establish a characteristic reference
for each sgecies.
The structural and enVirOTInlental characteristics
(if Camptosorus rhizophyllus (L.) Link. make it s(lmephat
difficult to find but
interestin::~
to observe
~'lhen
fe,und.
It is a 't'7alking evergreen fern \Vi th the narrOH tapering
leaves radiating from a r')otstock.
grc und, leaf tips sprout ne'tV' plants.
1
Upon tDuching the
Wagner re!,)orts it
to be closely related to Asplenium sibiricum of ne'rtheast Asia. (7)
C.rhizophyllus :;roHs best
the shaded,
~~oss-gro\'1n
faces ,:,f
exposure.
Those specimens observed in Parke Co. Here
fund
am~,1Ug
moist
c~)ol
reported to
limest,~ne
~;n
cliffs Hith a northern
limest0ne :)utcroI'pin;2:s, deep ravines, and
noss covered
ran::~e
b;~nks.
Populations have
b~en
from Alabama and Georgia north to
lfinnesota and Uuebec as Fell as I'lest to Nissouri. (16)
16
Leaves are narrmv, triangular tapering \.-i th a ccrdate
base avera:Ling t\lel ve inches long to smaller basal leaves
of one inch.
The blades average seven inches long, 5/8
inch ,;·ri.cle; the periole abilut l-~- inches long.(ll)
plants gro\-7 flat upon the
-:~round
semi-erect with arching le.aves.
l'lavy to indented.
Youn~~
while older plants are
Leaf margins vary from
Veins are freely areol:,te or netted.
The rootstock is vertical vri. th bro,m evenly
spacl~d
scales.
The stalk is cU3rk brmm, short and flattened; it may be
smooth green above., scaly at the base.
Brc\·m sori c:.re
scattered from the base to tip on the leaf undersurface.
The indusium is inconspicu0us.(4)
See Figures I, IV.
According to the data collected for Camptosorus
~:'hizophyllus
(Table IA) the average total height of the leaf
FaS
15.6 em
for hlenty-five specimens from parke County, Indiana.
The average frond length 17.1 cm; the average stem length
2.9cm; and the mean of frond Fidth 1.3 cm.
An analysis
of the leaf length variation for C. rhizophyllus may be
f-;lund in Table lB.
Asplenium platyneuron is ·)ne of the most i'liclely distributed spleel1"t;J()rts.
Populations rEm;:::;e from Florida "":est
to Texas, Kansas to Visc:msin, throughout the Appalachians
and northeast.(16)
It ,3rm·lS in shaded i;-oods, fields, and
banks Hhere rocky soil is ,·;rell drained and Goist.
Those
sDecimens observed for this study ,vere collected in I]reene
17
(c.~
.
~3 em
C(JtnpfosorU6 r- h',:zophylllt s(L)Unk
R.rke Co.) IndlQhQ
'1-IS-~5"
San ds-rc he cu+c..r-o pp i n3
18
Figure 4
--Nature Sporophyte of
Camptosorus rhizophy11us (L.) Link
Parke Co., Indiana
7-18-65
19
Table I B
Analysis of Leaf Variation am:.:Jng Camptosorus
;:hizophy11us (L.) ):..irik
Collection
S1te
Parke Co.
Leaf
~-le.ight
8.0
19.5
.28.5
27.2
';.7
14.0
17.5
26.0
18.5
17.1
11.0
17.1
9.5
23.0
1":-.7
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
11.3
9.5
:) .8
6.2
17.7
:~~! .5
4.9
6.0
1 1::-.5
21.4 em
Total:
Jr:ean:
3~'2.1
-
em
15.7 em
l)eviation
=-'"
De.viaticm2
-7.7
3.8
1;~.8
11.5
-6.0
-1.7
1.8
10.3
2.8
1.4
-3.3
1.4
-6.?
7.3
-1.0
-4.4
-6.2
-6.9
-9.5
2.0
L>.8
-10.8
-9.7
-1.2
5.7
--
5;', .5
14.5
166.5
132.8
36~O'
2.8
3.2
106.1
7.8
1.9
10.2
1.9
38.5
53.5
1.0
19.6
38.5
47.9
:.0.2
l}.O
190.1
110.7
94.1
1.4
32.5
,--1255.46 em
Varienee: 5G.14 em
.standard
Deviatiun:
7.13 em
21
12.5<:.m
Asplenium plat-tDeuron (L)Ookes
Monrae
Co.) .1.ndiana
7-25-~5
I"f'\ois+ leaf compost
22
Figure 5
~Bture
Sporophyte of
Asplenium platyneuron (L.) Oal:es
Nonroe Co., Indiana
7-25-65
23
The av(?.rof.Se fr::lUd 12.nsth ,las 17.1 cm; the
ave.ra~;e
len:;th 3.2 cm; and the mean of frond \\riclth 2.6 em.
analysis of the leaf heii2;ht variotion in
~:l..
s'cem
An
)latyneuron
f:lay be f c,und in Table 2B.
Asplenium ebenoides
(:.~cott)
Wherry or :-[
sorus '2benoides is a small tufted variable
f(~l~n.
"~spleno­
ever~;reen
It is Sup,)osed by seme 'i7orkers to be the earliest
established fern hybrid; and may occassionally be :c'.und
f:2rtile.
A 'vide distribution of A. ebcnoides includes
an area of Verm,mt to lIissouri and into Alabama.
~~)op­
ulati,ns are re90rted in Noo'tv York, H,,:]ssachusetts, Gonnecticutt, :eennsylvania, New Jersey, liaryland, Illinois t
Virginia and Indiana.(4)
It
:rous best upon lir::test;'Ue
'utcrolnin;:,ss and cliffs \There the sc,il is moist sand or
rich loam in a north(~rn exposure.
!.eaves of ~. e.be.noides
normally do not exceed 5 - 6 inches, but may be up to
12 inches lont;.
at the apex.
They are uider at the base and tal)ering
Leaf Sha!?e is extremely variable ':Jhereas
the lO\·,rer 1/3 leaflets may be cut ';?artially or E~ntirely
t·:) the axis; I,rhile the u;,)er ';:/3 leaflets may bE~ fused
ryartially or scalloped
'Iiri
th fe,·, indentations or straight
margined up to the apex.
fiost leaves grm·: flat upon the
ground 'i'lith longer leaves
~ointins
appard.
are freely forked and rarely netted.
Leaflet veins
Sari are linear
24
Table II B
Analysis of
Collection
Site
Greene Co.
11
!T
11
f1
110nroe Co.
!l
Green~
Co.
!T
If
"
Honroe Co.
Greene Co.
tt
Honroe
Greene
Nonroe
Greene
Honroe
Hlinroe
T.:)tal~
Nean':
Co.
Co.
Co.
Co.
Co.
Co.
L-~af
Variation
amon~
Leaf Height
16.5
28.8
31.7
32.0
19.5
12.0
15.0
29.0
25.7
17.0
17.9
15.0
16.0
19.5
12.5
25.2
20.8
22.8
13.7
15.6
em
em
em
em
em
em
em
em
em
em
em
em
em
eql
em
em
em
em
em
em
405.8 em
70.3 em
Asplenium, ~atyneuron
( L.) () .es
Deviation
2
'Deviation
-3.8
8.5
11.4
11.7
-0.8
-8.3
-5.3
8.7
5~4
-3.3
-2.8
-5.3
-4.3
-0.8
-7.8
4.9
0;5
2.5
-6.6
-4.7
14.5
72.0
130.0
135.0
6.4
69.0
28.0
73.0
29.0
10.9
7.9
28.2
18.5
6.4
Gl.4
24.0
2~5
6.2
':~3.5
22.9
775.6 em
Variance: 38.8
Standard
Deviation:
6.3 em
25
th(~y
shaned being nearer the midvein than mRrgin;
frequently d;,;uble.d as ,Jell.
are
The indusium is silvery
and found at an angle, or to (me side of the sorus.
The leaf axis is brown beloH; green c:bove.
A short
brittle shining purple-brm'm str:lk projects .Erom the
short vertica1. dark rootstock.
The roots are black,
v]iry, and shallow-creeping. (11)
See Figures III, VI.
Externally, many of these characteristics resemble those
of both
c.
rhizophyllus, A. platyneuron.
Those specimens
observed of A. e.benoides (Table. 3A) borrol·red from the
Indiana University Herbarium and the Chicago Nai:ural
~~Iistory
lfuseum varied "\,Jidely from locale to locale in
over-all
me.asurem'~nt.
(Tabl~.
30)
The mean of all
area measurements for total leaf height HaS 10.1 cm
for tFenty-one specimens.
all areas
~7as
The mean frond length for
7.5 cm; the mean stem length HaS 2.5 cm;
and the mean frond vTidth "7as 2.8 cm.
An analysis of
the leaf height variation in A. ebenoides may
in Table 3B.
bE~
fe,und
26
/5".5 c...rn
benoides
Aspleno$orc.lS (ScottI
Wherry
5-1~-",q
'um
.:c. U. Herbar~,fo,
60315
27
Figure 6
.
I
}~ture
Sporophyte of
j.o,.sp1enosorus ebenoides (Scott)Uherry
Nonroe Co., Indiana
5-14-49
28
Table III B
Analysis of Leaf Variation among Astlenium ebenoides
Scott)
Ufierry
Co11ecti-.)n
site
Slosson hybrid
If
If
11
"
"
II
IT
II
,~
It
'rotal
l'1ean
Leaf Height
10.3
9.6
11.7
11.8
11.9
15.5
12.3
13.3
11.0
8.0
9.4
124.8 cm
11.3 em
Deviation
Deviation2
-1.0
-1.7
0.4
0.5
0.6
4.2
1.0
2.0
-0.3
-3.3
-1.9
Total
Variance
Standard
Deviation
1.0
2.9
0.2
0.2
0.4
17.7
l~O
4.0
0.9
10.9
3.6
42.7 em
3.9 cm
1.9 cm
29
'fable III
C
Analysis of leaf Variation among ASElenium ebenoides
(Scott)
\ Jherry
Collection
Site
Honroe Co.,
Indiana
nOlo
hl m1.ng t on,
T)ela~!are
II
II
College Hill,
3aston, Pa.
II
"
Havana Glen,
Hale Co. , A.la.
If
II
II
II
"
II
If
"
Total
l-1ean
t.e.af Height
em
em
em
em
em
em
em
em
em
1l~.8 em
15.0 em
14.0 em
7.0 em
12.0 em
5.5 em
6.4 em
14.7 em
11.8 em
11.6 em
13.5 em
201.1 em
10.05 era
15.5
18.0
3.4
7.3
3.6
6.6
5.0
4.2
10.7
Deviation
5.5
8.0
-6.6
-2.7
-6.4
-3.4
-5.0
-5.8
0.7
4.8
5.0
L~. 0
-3.0
2.5
-4.5
-3.6
4.7
1.8
1.6
5.5
Deviation2
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
30.2
64.0
4-3.9
7.2
l~l.l
11.6
:::5.0
33.9
4.9
23.2
;~5. 0
16.0
').0
6.2
20.4
1.:>.0
22.0
3.2
2.6
12.3
~otal 4;)9.3
Varianee. 20.5
3tandard
Deviation
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
em
4.5 em
30
Spore Norphology
As indicated by NcVaugh in his article (8) the epispore is present in both the C. rhizophyllus and A. platy
neuron.
It may appear as pro jecting c ,ntinuous "rin:.:;s or
rid'>:es on the spores; and it is not loosely attached.
Hrinl<.les of the epispore may also form ridges \'lhich become.
continuous.
In C. rhiznphyllus folds of the epispore
appeared to form a \"ide c·.:-nspicuous '\dng cc)mpletely abuut
the light to dark bro'lm spore.
vary fr·:)m 25-30 X ?7-35 mu.
Spore size is reported to
H01;vever,
in~.
,)latyneuron
the epispore rid::;es tended to anast·:)mose and 'Here thin,
not ro'tmded or obtuse.
The light or dt:1rk brO't·m 8)OreS
vary 30-40 X 35-45 mu. from reports.
The sori Gf C.
rhizophyllus were irregularly scattered into oblique or
paired grnups from the leaf base to tip.
Straight, elon-
gate sori of A. platyneuron 'vere regularly paired upon
veins branching from °che midvein.
(.~ori
were spaced in pairs beinG closer to the
leaf base and
m1en
;~Jne
tap~ring
of A. ebenoides
mar~~in
at the
touard nidvein at the leaf apex.
exarlines the structure of a spore three
main portions may be found; the protoplast is the innernost 1iving portion enc.'lsed by a thin cellulose layer or
intine; ,snd the. exine ·t-7hich is an outer cuticularized
very resistent layer. (3)
31
Based upon the measurements of simple slide prep...
arations view'ed pith an oil immersion lens and an ocular
micrnmeter, an analysis ':las made of the variation J_n spore
size af:l)ng several specimens of the collected material and
that borroued from hm herbariums.
Calculati_ns of spore
length and width were made ::::rorn recorded measurements ;:ihick
were used to obtain a mean, deviation, variance, and a
standard deviation of the
me~suren2.nt8
~\'t
recorded.
least
25 different s')ores 'Pere measured for each fern species.
Calculati ,ns of spores i=rorn,"::;. rhizophyllus revealed a
mean of 24.6 X 31.9 mu. (Tables IVa, IVb) C8 1cu]_ations
fr(:El snores of A. r,latyneuron revealed a mean of ::24.6 X
32.9 mu. (Tables Va, Vb)
lfeasurem'2.nts of
~
eben·)ides
include s;:>ores from the L:.nroe Co. 8rea ,nly.
S'.3 Z 11.3
TaU.
\',a8 revealed. (Tables VIa, V1b)
1-">.
r..1can of
Referring
back to each Table re.s;;ecti vely, it ap?2.2red that
c.
rhizo-
phyllus shO\·!ed the greatest variability in Ileasurement \lith
a standard deviation of 5.7 mu in 't,lidth and 6.3 mll. in spore
len:~th.
Spore
measurem~nts
of
plat~euron
sho\'Je.d a
stcmdard deviati 'n of 4.9 mu in spore . .·:ridth and 5.2 mu in
spore length.
The standard deviation of A. eben,.)ides
appe::1red 10\·!est Fith a 3.4 mu ,-!idth and 3.5 mu len;:::;th as
compared -to its average measurements.
'I'he chanE:e of
envir0nment mi~sht be consicl2.red an·i.nfluential source in the
..32
.1' ab1e
IV A
Spore Size Analysis of Camptosorus rhizophy11us
(L.)
Spore length
34 mu
20 mu
26 mu
20 mu
37 mu
34 "'1U
40 mu
26 mu
31 fiU
23 rou
37 mu
!+O
fiU
34 fiU
31 mu
L~3 mu
L:·O mu
31 mu
.54 mu
43 mu
31 mu
28 mu
28 mu
::::6 mu
37 mu
26 mu
34 mu
28 mu
Total 862 fiU
He. an
31.9 rnu
Link
Deviation
2.1 rou
-11.9 mu
-5.9 rou
-11.9 mu
5.1 mu
2.1 rou
8.1 fiU
-5.9 mu
-0.9 mu
-8.9 fiU
5.1 mu
8.1
2.1
-0.9
11.1
8.1
-0.9
mu
mu
11.1
-0.9
-3.9
-3.9
-5.9
5.1
-5.9
2.1
-3.9
mu
mu
mu
mu
mu
mu
mu
::::.1 mu
mu
mu
mu
mu
mu
mu
Varinnce
4.4 mu
141.6 mu
~·A.9 mu
141.6 mu
26.1 mu
4.4 mu
65.6 mu
34.9 mu
0.8 mu
79.2 mu
26.1 rou
65.6 mu
L~.4 fiU
0.8
123.2
G5.6
0.8
4.4
123.2
0.8
15.2
l5.2
mu
mu
mu
mu
mu
rou
mu
mu
mu
3l~.9
mu
26.9
34.9
4.4
_15.2
1074.3
mu
mu
mu
39.8
fiU
~
mu
::,tandard
')eviation
6.3 mu
33
Table IV B
·Spore Size Analysis of :Jamptosorus
~L. j
Spore T'idth
28 mu
lC~
23
14
23
28
28
20
20
14
20
.......j ' i
.<':..)
23
26
:·1
28
rou
rou
rou
mu
mu
mu
mu
rou
mu
mu
mu
mu
mu
mu
mu
:.0 rlU
31 mu
28 mu
26 rou
31 roil
31 rou
/-.0 mu
31 mu
17 mu
31 rou
23 mu
---Total 665 nu
~n
rhizoEh~llus
• i,
,_,llli-_
Deviation2
DeviatL,n
.5.4 mu
-10.6 mu
-1 .• 6 mu
~10.6 rou
-1.6 rou
3.4 mu
3.4 mu
-l~.6 mu
-4.6 mu
-10.6 rou
-10.6 fiU
3.4 mu
-1.6 mu
1.4 mu
6.4 mu
3.4 rou
-4.6 mu
6.4 mu
3.4 rou
1.L~ mu
6.4 rou
6.4 mu
2i.4 mu
G.4 rou
7.6 mu
6.4 mu
-1.6 mu
Total
Variance
11.6
112.3
2.6
112.3
2.6
11.6
11.6
21.3
21.3
112.3
112.3
ll.6
2.6
lUU
mu
r:1U
mu
mu
mu
l:lU
lUU
mu
mu
mu
mu
mu
rou
rou
mu
mu
1.9
41.0
11.6
21.3
[}1.0 I!lU
11.6 mu
1.9 lUU
[tl.O lUU
[~1.0
lUU
lUU
11.6
Ltl.O rou
~)7 .7
fiU
[tl.O rou
2.6 rou
9l1.9 mu
33.8 lUU
Standard
Deviation
5.7
lUU
34
Table VA
Spore Size .l..na1ysis of Asplenium p1atyneur',\U (L. ) C2kes
Spore l.e.nsth
Deviation
;;eviation2
34 mu
31 mu
.) I mu
34 fiU
20 fiU
30 fiU
37 mu
1.1 fiU
-1.9 mu
4.1 mu
1.1 mu
-1~.9 mu
-2.9 mu
~r .1 mu
1.1 mu
-1.9 mu
7.1 mu
-4.9 mu
-6.9 mu
-4.9 mu
4.1 mu
7.1 mu
1.1 fiU
-1.9 mu
-4.9 Llll
4.1 mu
7.1 mu
-4.9 mu
1.1 mu
10.1 fiU
-4.9 mu
1.1 mu
1.2 mu
.3.6 mu
16.8 mu
1.2 fiU
166.4 fiU
.].4 fiU
16.8 mu
1.2 mu
~...,
3l, fiU
31 mu
40 mu
28 mu
26 fiU
LO mu
37 fiU
L}O mu
3l~ mu
.31 mu
28 fiU
~n
37 mu
,,~O mu
28 mu
3l~
mu
l .. .:; fiU
28 fiU
mu
Tctal 824 mu
3'~
3.6 mu
50.4 mu
Total
24.0 mu
47.6 mu
24.0 mu
16.8 mu
50.4 fiU
1.2 mu
3.6 mu
24.0 mu
16.8 mu
50.4 mu
24.0 mu
l.2 mu
102.0 mu
~4.0 mu
1.2 mu
680.8 mu
Variance
27.2 mu
:3tandElrd
Deviation
5.2 mu
35
Table V B
~:;pore
Size Imalysis of Asplenium platyneuron (L.) Oakes
Spore
~!idth
31 mu
22 mu
27 mu
21 flU
10 mu
20 mu
28 mu
28 fiU
28 mu
28 mu
23 mu
20 mu
23 mu
34 rou
26 mu
26 mu
20 mu
23 mu
26 "1U
26 BU
17 mu
28 rou
31 rou
23 mu
26 mu
Total: 615 rou
Hean:
24.6 mu
Deviation
Deviation2
6.4 rou
-2.6 r·1U
2.4 mu
-3.L~ mu
-14.6
41.0 mu
6.7
5.8
12.9
214.2
21.2
11.6
11.6
11.6
11.6
2.6
21.2
2.6
88.3
1.9
1.9
21.2
2.6
1.9
1.9
fiU
4.6 mu
3.4 mu
3.4 rou
3.4 fiU
3.4 fiU
-1.6 rou
-4.6 rou
-1.6
IllU
9~4
mu
mu
mu
mu
mu
mu
1.4
1.4
-4.6
-1.6
1.4
1.4 rou
-6.4 fiU
3.4 rou
6.4 mu
-1.6
41.0
fiU
lllU
fiU
mu
pm
rou
mu
fiU
mu
mu
fiU
mu
fiU
mu
rou
au
fiU
E1U
mu
BU
11.6 fiU
41.0 rou
2.6 rou
1.9 mu
fiU
1.4 mu
Total:
variance:
Standard
Deviation:
592.4 mu
23.7 rou
4.9
fiU
36
Table
vt A
Spore Size Analysis of Asplenosorus ebenoides(Scott) Uherry
Spore Length
20 mu
10 mu
15 mu
16 fiU
8 rau
14 mu
16 mu
7 mu
11 mu
12 mu
7 mu
5 mu
6 L1U
14 rau
8 FlU
11 mu
17 mu
11,mu
14 mu
20 mu
8 rau
14 mu
20 mu
14 mu
10 rou
Total:
Hean:
282
fiU
11.3 mu
•
•
2
Dev~at~on
Deviation
9.8
-1.3
3.8
4.8
-3.3
2.8
-1.3
-4.3
-0.3
mu
mu
fiU
mu
rau
0.7
18.4
39.5
27.[:
7.9
10.8
0.1
34.0
0.1
7.9
77.9
10.8
7.9
77.9
7.9
1.6
fiU
./
uu
fiU
mu
fiU
mu
rnu
ElU
rau
O.l mu
.".
-::>.:.> fiU
2.3
-3.3
-0.3
5.8
-0.3
2.8
fiU
fiU
.
Tn1..'\.
mu
0.8 mu
-4.3 mu
-6.3 fiU
~
96.4
1.6
14.6
23.3
10.8
7 0
1.6
18.4
mu
mu
fiU
mu
mu
mu
8.8 mu
-3.3 fiU
2.8 rau
8.8 rou
2.8 mu
-1.3 mu
rou
rau
1:1.U
mu
mu
Iau
mu
mu
rau
fiU
fiU
mu
fiU
rau
mu
Total:
315.9 mu
Variance:
12.6 mu
Standard
Deviation: 3.5 mu
37
Table VI B
Spore Size Analysis of Asplenosorus ebenoides (Scott) Hherry
Spore Hidth
17
7
13
13
7
12
mu
mu
mu
mu
mu
mu
8 fiU
6 E1U
10 mu
11 mu
6 r.:lU
.3 fill.
6
8
6
8
14
mu
mu
mu
fiU
mu
6 BU
11
14
6
11
11
12
6
Total:
Mean:
mu
mu
fiU
rlU
mu
mu
mu
232 mu
9.3 mu
Deviation
Deviation 2
8.8 mu
2.3 mu
3.8 mu
n
mu
-'.0
2.3 mu
2.8 mu
1.3 mu
77.9 mu
5.3 mu
1 L}.6 fiU
1L~. 6 mu
fiU
.5.3
7.9
1.6
10.8
1.6
mu
mu
mu
mu
mu
mu
mu
L:-.8 mu
3.3 mu
1.3 fiU
2.8 mu
3.3 mu
1.8 mu
1.8 fiU
2.8 mu
3.3 mu
10.8
59.5
10.u
0.7
10.8
1.6
23.3
10.8
3.4
7.9
10.3
3.4
3.4
7.9
10.8
3.3 InU
1.3
1.8
3.3
6.3
3.3
0.8
3.3
1.3
fiU
mu
mu
mu
rau
3 .L~ fiU
Total:
Variance:
Standard
Deviation:
mu
fiU
r.:lU
mu
mu
fiU
rou
mu
mu
fiU
mu
mu
BU
mu
mu
293.9 mu
11.9
FlU
3.4 mu
38
variabili ty of the form.er two fern species; f8. rticularly
those of A. platyneuron.
In the event that severol plants
of each fern species H·:)uld have been in varying stages of
maturity, which is probable, the spore size measurements
'Hould
t~l.en
bot be
repl~esentative
of a given populaticn
in a specific sta.ge of development.
39
DISCUSSION
It ,'las observed from studies of
Camptosor~
rhiz()phyllus and .i'.I.splenium platyneuron that both species
as '!Jell as their reported hybrid tend to prefe.r a habitat
\;n.th a northern exposure that is similarly moist, shaded
and ,..yell draine.d.
All fhree fe.rns reqtlire a soil 1;nth a
high lime content 2nd is-:;lated location.
fically different, Pagne.r
~)oints
forms of A.• platyneuron and
£.:.
,out
t~~at
Although supe.rthe tjTpical
rhizophyllus con·tain 36
pairs (2n=72) of chrnmosome.s.(16)
During the yroduction
of spores, the ::-tairs separate intc) individual nigrating
chromosomes \'lhich ultimately form ne\<! ST)Ore. cells 'I;.n.th
36 chr'.,mosomes e.ach.
The typical il.. ebenoides ,a hybrid
of the tHO species (Slosson 1902), ·.-.iffer::; by b.2ving its
bvo sets of chromosomc::s c:)ming from
~)are.nts
,:hich are
genetically distant.
During spore production chromosomes
do not !:=lair normally;
bu~:
Fligrating c:1romosomcs.
remain 72 sin.s;le irre.Gularly
Since the resul tins: s)ores \Jill
not receive balanced chromosome. sets, they ,bort be.cc,ming
sterile.
For li. ·2be.noides to become fe.rtile , it Dust
have d.;uble. the number of cLromosomes expected (4n=144).
\fnen this occurs, it is possible for each original parental
set t:) h"ve a }i.Enlog2us set ui ttl 72 pairs.
In the chronato-
graphic study of Smith and t:evin (1963), conducted for the
40
purl?ose of taxa identification, [>,enomc designatic.lUs of
the Aspleniums 'pereiven thus:
Asnlenium
rhizophyllum
•
2n
iUt
ASDlenium platyneuron
2n
2P
x ebenoides
2n
::':'It
4n
PPRR (13' )
t
l~"sDlenium
e
.
As")lenium eben Jides
Duri.n::~
their biochemical studie.s of hybridiz,'ltion Sruith
and Levine :cuund detectable substances of L"e p,1rcmtal
species Hithin ';;he intcrs:)ecific hybrid.
using observations of the. E10rphologict.l charactertistics as a cmparisnn 'if
t~:le thr'~e
ferns, the distinct
di:eferentiation of their structural c"mposition
evident.
~'7aS
The A. ,':latynenron iJaS zencrally a larc;er )leut;
its leaves were divided into alternate leaflets; the leaves
were erect to :;:e':0.i-crcct.
The periole color.Ln..: changed
from green in the young pVmt '1:0 chestnut bro,·m orlurpleblack in the. mature specimen;.
~
rldzophyllus, on the
other hand, ,ras :.{e.ncrally slishtly snalle.r in
\.7hole leaf
grounc1 •
~x)ssibly
t.·uchin~; .~r~und,
'-I:lhe le:3£ is
(~ntire,
·~n
both '::ides.
~bereas
s"ne
Ti1'1r~~in
\7:.l.vy or ::calloped
r):c
.c~nd
,·las light to dClrk bro'l;ffi only )CTt
T.ringed
the
curved and serc1i-erect to flat u:}on the
::iost of frond tips \Jere
s.iruuting.
11.(~i~ht;
8~ri
the way t
of the
narrOlvly
platyneuron
41
~'7ere
s:).?ced
ev·?-n1~7 o~»osite
and close. to the. uidve.in of
le.afle.ts; ·che. sori of C. rhizophy"'llus ,qere.
~(':und
scatte.re.d aL:mg the. marginal edges of lenflets.
of A.•
ebenoide.~
une.venly
Specimens
see.me.d to cc;mbine the. characteristics of
che.stnut to !)urple
p~tiole
of size 2nd leaflet form.
color and the variability both
j'l.pproximately 2/3 or t;le Zrond
h·9d separate. leaflets op;Jositely arranged; the remainder
being an entire fusion ;)f tb.e leafle.ts ui th a scalloped
or "mvy margin.
"hole le.af.
enclosed
1'he I!etiole composed only about
± of
the
The remainder of the pe.tiole extension "\'as
b~T ';;~Le
fused leaflets.
Jori Here both marsinal
and close to the midrib portion oE the. leafle.ts.
leaflets appeared fan-shaped
e.nds of radiating veins.
~:i th
Some
sori at trle marginal
Since. :c;ucce.ssful cultures of
the. mqture :z;ameto}hytes -';7ere n·::>t ob·taine.d in this study t
c')m:')arisons i'Jere :llade. sole.y "m the. r.1orphological
stru.cture.
The. three. ferns eX8111ine.d "lere found to possess
trilete spores.
Tlvmc~h
greatly variable in size. and
detaile.d structure, the ;:;e.ne.ral
~latte.::..~n
c>f the. sqores
'[las found to be. similar. (See Figure.s VII thr,:,.u[;h IX)
The. e.pis:}ore.
borde.r
u~on
a~)I)e.are.d
in v3ried uings or as an
the spores.
1.:me.v~n
lIany ridges \Jere found t .. be.
continuous Hrinlde.s in S3)ore.s of C.
A.. I'latyneuron, and A. e.be.noide.s.
rhizophyll:..~,
42
Figure 7
Spore Ylaterial
Camptosorus rhizophyllus
t
IO.-y
-I
43
Figure 8
Spore l'1aterial
Asplenium platyneuron
44
Figure 9
Spoee lla.terial
Asplenosorus ebenoides
45
In ~ rhizophyllus the \'iide ring of folds a!)peared completely to surr:Juncl the spore; uhereas in 1i...
nlat'me'lU.~on
+
,...
the s))ore 'pings \,rere broken :)eing thinner or nen-existent
on <.)ne side or the
oth(,~r.
Both ferns
)ossess(~d
spores
Ilhose ridges -';Tere anastomosing but }:,ose of A. ;)latyneuron
\Vere ,Clore nume.rous and thin; not r':illnded or ob;:use as
those of
.2..!.
rhizo~hyllus.
The s:)ores of sl>ecimens collected
fc)r -this study ranged 17-34 X 26-40 mu for
?latyneuron
and 14-31 X 21-37 mu for C. rhizophyllus.
the isolAted
e}~tremes,
uniform in size.
A.
and
~benJides
.::~;en2.ral
mos-;: of the
s~)ores
In soite of
examined uere
Th'Jse spores examined from specimens of
, hOv1ever, T1ere-reatly variable b· ·th
appearance.
size
c2he ,7in::ed e.pis)ore seemed both
widely c:nspicuous and thinly c::mtinu::ms.
s~)ores
iL~
~ddge.s
upon the
",ere both finely netted cmd variously curved.
avera~~e
size ivas 9
~{
The
11 mu.
Using the structural characteristics of external
nl;:mt form :C1nd spore structure for comparison, it \71;uld
appear that A. eben.,ides seems to extr.=:Tnally c:.:;mbine the
most noticeable char?cteristics of
A. platyneurcn.
.2..!.
rhizophyllus and
A chromosome c(,unt of material from the
parent ferns ,ruld have been a
[,lOre
useful comparison as
"lOuld a chrc:·mos-rne count from A.
~benoideG.
A.
--
~)lausible
eben:")ides does anne.ar to be. a
ferns A.
~.
platyneur~
and
......
t~i
rhizppl:1yllus.
Superficially
hybrid of the
46
!)urin:.; the approxim:3te period cnver:rng June, 1965
thr,:-:·ughtFebru;~ry,
1966 a study has be.en made of C. rhizo-
phyllus (L.) Link and A. platyneuron (L.) Ogkes in order
to learn about th ::ir possible troonomic relationship Gnd
the significance of their spore morphology.,
been c')lle.cted from the dried material; simple slide
makin~
techniques learned; and microscopic measurements
observed.
The strong possibility Gf a valid hybrid,
Asplenium ebenc)ides (:::cott) ';!herry, bebdeen "che two fern
species has also been observed and found to be supported
by external structure c(lmparisons of the conspicuous
characteristics present.
In order to form
Q
definite
conclusion ab()ut the taxonomic relationship of
c.
rhizo-
phyllus 2nd A. platyneuron, it T,7ould be. necessary to
make other studies of a direct nature.
If,ine
,,~'ished
to learn more about the tax, 'nomic relationships and
hybrid nature of the A,spleniumrnu}?ing, studies of
chromosome counts or biDchemical constituents should be
attempted.
'lne major
obst~.cle
invf)lve.d in chr:.:,mosome.
studies \7ould be a succ(:;ssful culture of the spore
material 'Nhereby mature gametophytes ,,,ould be available.
47
BIBLIOGRAPHY
1.
Bold, Harold C. 1957. Norphology of Plants.
Harper and Brothers. Neu York, li. Y.
2.
Britton t
Ferns l l •
Donald 11. 1953. "Chromosone :~tudies on
Journal ~f Botany. 40:575-583.
Arneric~n
3.
Bro'Wn, Clair li. 1960. tfTi!hat is the ~lole of $yores
in Fern TaxonoP1.y?1I Arrrerican ~ Journal. 50:6-14.
4.
Cobb, Bou:3:~1ton. 1956. <s~ Field Guide '.i~0 The Ferns
And Their Related Families. Peterson Field Guide
Series #10. Houz:;hton l1ufE"1in Co. Boston, ljass.
5.
Erdtman, /}. 1943. An Introduction To Pollen Analysis.
The Ronald Press Go. He't-l York, n. Y.
6.
Gray, Peb~r. 1964. Handbook of Basic I:.icrotechnique. Nc;3-r€:l'tV'-Hi 11 Book Co. 3rd ~~cli t • He'l:'] York, N. Y.
7.
Haxon, t']i11iam R.
1918. fI,A Hew Hybrid L'l.splenium" •
.:.Jnerican Fern J'ournal. 8:1-3. January-Narch.
8.
l!cVaugh, Roge.rs. 1935. "Studies on the Spores of
some Hortheastern Ferns. It ."-_rnerican Fern Journal.
25:73-85. July-September.
9.
Hiller, David F. ,Jnd Glenn '). Blaydes. 1938.
11athods and Haterials for Teacl1.in;; Biological Sciences •
llcGraw-Hlll Book Co. Ne.vl York, 11. Y.
10.
Sass, John~. 1958. Botanical Hicrotechnique. 3rd Edit.
IO\va State. Colle:;e J.'ress. L':;.mes, 100'1a.
11.
Shaver, Jesse L. 1954. Ferns of Tennessee ~jith
Their Fern :\.llies ::.:xcludeCf. ·}eorge Peabody College for
Teachers. rlc:tshv:tlle, Tennessee
12.
Slosson, Nargare1;
1902. "'l'he Origin 0:1: Asplenium
ebenoides. tI
BuL.. etin of the Torrey Botanical Club.
29:li87-4~)5.
i1.U~ust.
48
13.
Smith, Dale H.
and Donald li. Levin. 1963. itA
Study of rteticulate Z:volution In
The Ap)alachian :.s:)lenium Complex. It
il.LK~rican
Journal of Botany. 50:932-950. October.
Chrol~lato0raphic
14.
')tebbins, G. L.
1950. variation and
in ?L:mts.
Ne~'7 York,
P. Y.
15.
~la~;ner,
r~volution
~,;arren ~:.
1953. TtA ,.jytological Study of
Jche . ApP·llac ::,ian Spleen;7orts lf •
~.2ulerican Fern Journal.
43:109-11L~.
16.
• 1954. '!Reticulate Evolutim In
The i:..?palachian ;~spleniums. Il:i;volution. 8: 103-118.
J}.ll1.e.
17.
• and Kathryn Boydston. 1958. If A
Nevi l-Iybrid Spleem'70rt from ~~tificial Cultures at
Fern-Hood and Its Relationship to a ?ecul.iar Plant
:Ero;~'.. 'l:est Vir:;inia".
;>T:lerican 2e.rn Journal.
48: 145-159. '-
L:).
• and Robert :.:;. 'ilhi tuire.. 1957 •
nSpontaneous FrocL11.ction of )~ lIorpholo~c::ically Distinct
Fertile ~:~lloJoly~)loid ?Jy i:. 3terile :Ji)loid of Asplenium
:;be.noides." Bulletin of the 'l'orrey 30 ;:anical Club.
84: 79-39. l~;:!rch.
J
.